Selaginella tenuissima Fée (1873: 98)

Valdespino, Iván A., 2015, Lectotypification of Selaginella tenuissima and S. papagaiensis (Selaginellaceae) with the description of S. monticola, Phytotaxa 233 (2), pp. 153-165 : 157-163

Selaginella tenuissima Fée (1873: 98) . ( Figs. 2–5 )

Type:— BRAZIL. Rio de Janeiro: Alto Macahé, [12 March 1870], Glaziou 4499 (lectotype: P [ P 00573924]-image!, here designated).

Selaginella papagaiensis Silveira, Bol. Commiss. Geogr. Geol. Minas Geraes 5: 125. 1898.

Type:— BRAZIL. Rio de Janeiro: Minas Geraes [Gerais], In campis humidis in Serra do Papagaio, Minas, November 1897, Silveira s.n. In Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2603 (lectotype: R! (as Herb. Silveira No. 154) [179579], here designated; isolectotypes: B [ B 20 0120913 (b)]-image!, BM! (as Herb. Silveira No. 154), P [ P 01242339 (lower, center gathering)]-image!).— Selaginella papagaiensis Silveira forma α [alpha], Bol. Commiss. Geogr. Geol. Minas Geraes 5: 126. 1898. Nom. superfluous for S. papagaiensis forma papagaiensis .

Selaginella papagaiensis Silveira forma β [beta], Bol. Commiss. Geogr. Geol. Minas Geraes 5: 126. 1898.

Type:— BRAZIL. Rio de Janeiro: Minas Geraes [Gerais], In campis humidis in Serra do Papagaio , November 1897, Silveira s.n. (lectotype: R! (as Herb. Silveira No. 155) [ R 179614 ], here designated ; isolectotypes: B (as Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2755) [ B 20 0120911 (c)]-image!, P (as Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2755) [ P01242339 (upper, center gathering)]-image !).

Habitat and distribution:— Selaginella tenuissima grows as terrestrial or epipetric plant at high elevations in Cerrado (tropical savannah) and Atlantic Rainforest vegetation. It is endemic to Brazil where it is reported from the states of Bahia, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, and São Paulo ( Alston 1936, Alston et al. 1981, Hirai & Prado 2000, Hirai 2015). Its distribution in Bahia is formally vouchered here.

Conservation status:—According to Hirai (2015), the threat to Selaginella tenuissima is considered of Least Concern (LC).

Specimens examined:— BRAZIL. Bahia: Morro do Chapéu, BR 052 , vicinity of bridge over Rio Ferro Doido, ca. 18 km E of Morro do Chapéu, 11°33’01”S, 41°09’21”W, 1000 m, 17 June 1981, Boom & Mori 1264 ( NY), Serra do Tombador , Rio Ferro Doido , ca. 18 km E of Morro do Chapéu, 1100 m, 21 February 1971, Irwin et al, 30741 ( NY) . Minas Gerais: Caparaó, Parque Nacional do Caparaó, along trail from Tronqueira to Terreirão , Along Rio José Pedro , 20°26’S, 41°44’W, 1970–2350 m, 31 October 1994, Buck 27064 ( NY), Alto Carapaó, road between Tronquera and foothill of Pico Bandeira, region of Três Lagoas, road to Pico do Cristal , 20°25’05”S, 41°48’56.4”W, 1800–2560 m, 23 November 2006, Salino et al. 11434 ( BHCB n.v., PMA), 20°27’S, 41°50’W, 1000–1970 m, 15 September 1984, Vital & Buck 11554 ( NY), Caparaó Novo, Cachoeira Bonita, 20°27’S, 41°50’W, 1600 m, 15 September 1984, Vital & Buck 11588 ( NY) ; Itabira, 1891, Damazio s.n. ( RB) ; Itaguaré, Passa Quatro , 9 May 1948, Brade & Silva 19040 ( RB) ; Ouro Preto, Serra do Espinhaço, lower slopes of Pico de Itacolomi , ca. 3 km S. of Ouro Preto , 20°23’15”S, 43°30’21”W, 1 February 1971, Irwin et al. 29551 ( NY) ; Parque Nacional do Itatiaia, vicinity of Brejo da Lapa , km 8 along entry road near border of Rio de Janeiro, ca. 22°22’S, 44°45’W, 2120 m, 5 July 1991, Vital & Buck 19697 ( NY, PMA) ; Serra do Papagaio, November 1897, Silveira s.n. ( B (as Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2755) [ B 20 0120911 (c)]-image, P (as Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2755) [ P01242339 (upper, center gathering)]-image, R (as Herb. Silveira No. 155) [179614] ); Serra do Picú , 1900 m, 1879, Schwacke 1640 ( RB), 1887, Schwacke 5897 ( RB) . Rio de Janeiro: Itatiaia, Parque Nacional do Itatiaia, Abrigo Rebouças , 2300 m, 11 October 1977, Maas & Martinelli 3172 ( RB), Vicinity of Itatiaya , 22°29’46”S, 44°33’47”W, 26–30 July 1915, Rose & Russel 20534 ( NY), Serra do Itatiaia , 2300 m, 1913, Brade 8626 ( R), Serra Itatiaya [Serra do Itatiaia], 2250 m, June 1913, Tamandaré & Brade 832 ( RB), Serra do Itatiaya , 2250 m, 4–10 June 1913, Tamandaré & Brade 6508 ( NY) ; Serra dos Orgãos, Rio Castellos , 19–20 February 1933, Brade 11532 ( R), 27 February 1933, Brade 12439 ( R) ; Terezópolis, Campo das Antas , 2100 m, 9 October 1929, Brade 9662 ( R) ; Mpio. de Resende, Itataía National Park, South face of Mt. Itatiáia , along path above “Macieriras”, 2060 m, 29 July 1966, Eiten & Eiten 7621 ( MO) ; October 1871, Glaziou 4499 ( B [ B 20 0095602] (1) & (2), C [ C10020554 ]-image, K [ K000589061 ], P [ P00044826 ]) . São Paulo: Campos do Jordão , 1 January1947, Leite 3673 ( MO) ; Jaraguá , 24 March 1912, Brade 5137 ( NY) .

Discussion:— Selaginella tenuissima is morphologically similar to S. monticola , which see for comparison. There is controversy regarding the correct typification of S. tenuissima as will be ascertained from the following discussion. According to Stafleu & Cowan (1976–1988: 821), Fée published his Cryptogames vasculaires (Fougères, Lycopodiacées, Hydropteridées, Equisetacées) du Brésil (Crypt. vasc. Brésil), in two parts. Part 1 was published in 1869 and Part 2 in 1872–1873 [1873]. Fée described S. tenuissima in Part 2 (1873: 98), citing a single collection “ Rio- Janeiro, Alto-Macahé, Glaziou 4499 ” without indicating a herbarium. Stafleu & Cowan (1976–1988: 818) indicated that Fée’s herbarium and types were originally at STR but are untraced and that Urban mentioned that Cosson acquired the Brazilian ferns, which are now housed at P. However, according to a further note by Stafleu & Cowan (1976–1988: 818) they alleged that Dodge saw Fée’s herbarium at RB in 1959. It seems that Dom Pedro II, Emperor of Brazil, bought Fée’s herbarium in 1871 ( Windisch 1982), apparently through negotiations with Glaziou, who at that time was Director of the Botanical Garden at Rio de Janeiro ( Maddi et al. 2013). Windisch (1982) further indicated that contrary to Stearn’s (1962) belief that Fée’s herbarium was lost at sea while en route to Brazil, it is now at RB. The types found at RB presumably include, among others, those of Fée’s pteridophyte species published in Part 1 (1869) of Crypt. vasc. Brésil. A search of digitized Selaginella specimens at RB ( JBRJ, 2015) yielded no “ Glaziou 4499 ” specimen there nor was it included among Fée’s fern types listed in that herbarium by Windisch (1982). This is to be expected given that S. tenuissima was published in 1873 and, thus, its type could not have been among specimens shipped to Brazil in 1871. The “ Glaziou 4499 ” collection is not recorded at STR by Hoff & Deluzarche (2013) either. Therefore, one may consider that the type of S. tenuissima would be among the specimens obtained by Cosson, however, these were apparently acquired before publication of the species concerned. A more plausible scenario is that Fée, himself, deposited the type of Selaginella tenuissima at P later on when he relocated from Strasbourg to Paris in 1870, as a result of the Franco-Prussian war ( Stearn 1962), three years prior to the publication of that species in his Crypt. vasc. Brésil. Part 2 in that city.

Among the Selaginella tenuissima holdings at P there are two specimens labeled “ Glaziou 4499 ”. One has a label from “ Herb. Glaziou Pl. de Rio de Janeiro, Selaginella tenuissima Fée, Cryptogames vasculaires du Brèsil (Suppl.) pag. 98, Tab. 108, fig. 1., N.4499. Alto Macahé, 12 Mars 1870 [12 March 1870],” P [P00573924]-image! ( Fig. 3 ). This specimen apparently is from Glaziou’s personal herbarium and was annotated by him, as the label and the handwriting on it match those depicted by Cremers & Boudrie (2013). The other specimen has a printed label that says “ Herb. Mus. Paris, Province de Brèsil, A. Glaziou ” with the collection number “ 4499 ” handwritten and a different imprint saying “ Reçu le 10 Janvier 187 6 [Received January 10, 1876]” ( Fig. 4 ). I examined this specimen (P! [P00044826]) and originally thought it to be the holotype, not being aware of the first mentioned. Neither of these two specimens, however, seem to have been annotated by Fée or to have his labels. They do not match his labels and handwriting, as shown by Cremers & Boudrie (2013) and Maddi et al. (2013).

Alston et al. (1981: 299) located the type of Selaginella tenuissima, Glaziou 4499 at C, in what could be construed as an unintentional lectotypification, while Hirai & Prado (2000) speculated that the holotype was at P with an isotype at C. The specimen at C [C10020554]-image! ( Fig. 5 ) has a printed label from “ Herbarium Eug. Warming ” with the name Selaginella tenuissima handwritten, “ Legit in Brasilia et communicavit A. Glaziou [collected in Brazil and shared by A. Glaziou]” printed and the collection number “ Glaziou 4499 ” handwritten in pencil, but does not indicate a collection date or a collection locality, as was given in the species’ protologue by Fée. Moreover, the handwriting on this label does not match either Glaziou’s or Fée’s.

Additional, presumable duplicates of “ Glaziou 4499 ” are located at B and K. The label on the specimen at K! [K000589061] specifies it was received on “10/71” [presumably October 1871], while the one at B! [B 20 0095602], does not indicate a collection date and is subdivided with the numbers “1” and “2” in pencil. The one sub-numbered “1” has a handwritten label with “ Glaziou 4499 ” and was annotated by Christ as “ Selaginella serpens [(Desv. ex Poir. 1814: 553)] Spring (1843: 228) ” and another separate determination slip by Hieronymus as S. tenuissima . The one sub-numbered “2” has a field tag label with the number “ 4499 ” attached to it, in what appears to be Glaziou’s handwriting, in addition to a label from “ Museum botanicum Berolinensis ” with the printed information “ Rio de Janeiro: A. Glaziou n. ” with the number “ 4499 ” handwritten and above it the handwritten expression “ Originalexemplar! ”. These two sub-numbered parts may in fact be a single gathering and do represent S. tenuissima . The specimen at K and the one sub-numbered “2” at B, I originally annotated as isotypes, but they do not indicate the collection locality information found in the species protologue either.

To complicate matters further, it is known that Glaziou deliberately tampered with his labels e.g., a) altered his collection localities, b) gave the same numbers to different species and even to different plant groups, c) placed his collection numbers on other collectors gatherings, and d) gave the same collection number to the same species collected in different periods ( Wurdack 1970, Cremers & Boudrie 2013, Da Costa-Lima et al. 2015 and references therein). This makes information on Glaziou’s collections unreliable or confusing ( Aranha Filho et al. 2007). Taking into consideration all of this and the fact that the information on the label in the specimen at C does not include the collection locality data as stated in the protologue of Selaginella tenuissima , the possible unintentional lectotypification by Alston et al. (1981) is here superseded according to Art. 9.19b of the International Code of Nomenclature, ICN ( McNeill et al. 2012). The specimens at B, K and one at P (i.e., [P00044826]) are equally excluded from type consideration because they do not explicitly indicate the collection locality as stated in the protologue and/or could have been gathered after 1873 as well. Consequently, it seems advisable to settle the nomenclatural identity of S. tenuissima by selecting as lectotype the one at P [P00573924] that best matches information in the protologue.

A synonym of Selaginella tenuissima is S. papagaiensis Silveira ( Alston 1936, Alston et al. 1981, Hirai & Prado 200, Hirai 2015), which also needs to be typified. Silveira (1898) described S. papagaiensis subdividing it in forma α [alpha], for which he cited his collection “ A. Silveira s.n. Nov. 1897 and H. Magalhães s.n., In Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2603 ” and forma β [beta] citing only “ A. Silveira s.n. Nov. 1897 ”. No specimen was specifically cited as the type of the species; thus as provided by the current ICN ( McNeill et al. 2012), a lectotype should be selected amongst the specimens cited for both forms. For this, I studied duplicates of the collections cited at B, BM, P, and R, all of which matched the protologue. The labels in the specimens at B and P agree better with the information in the protologue since they indicate “ Herb. Com. Geog. et Geol. Civitatis Minas Geraes No. 2603 ”, while those at BM and R, although with the same information in Silveira’s handwriting as the ones at B and P indicate they are from “ Herb. Silveira No. 154 ”. According to Oliver (2007) and Heringer (2012), Silveira worked as director of the “ Comissão Geográfica e Geológica de Minas Gerais ” (Comissão) from 1895–1898 and apparently his collections and those of H. Magalhães were deposited in the herbarium of that institution either as an institutional herbarium or as part of his own herbarium but were originally distributed under the institutional label. Nevertheless, upon finishing his directorship he apparently took his types and numbered them in a consecutive sequence as part of his personal herbarium as evident from later published species (e.g., Silveira 1908) and publication of Floralia Montium ( Silveira 1931) . His personal herbarium was acquired later on by R ( Oliver 2007). I studied all Silveira’s personal herbarium specimens thought to be the types of his Selaginella species during a trip to R in 1992, and later as part of a herbarium loan. In the case of S. papagaiensis , I physically compared the morphological characters and handwriting information of the specimens at R with those at BM and B, as well as with digitized images at P, against the protologues of the taxa here concerned. As a result, I am quite certain that the specimens at R must be the original material on which Silveira based his Selaginella species, including S. papagaiensis and the two forms. Therefore, in selecting a lectotype for these taxa I gave priority to those at R, albeit they have a different label as indicated above. I was also able to determine that H. Magalhães s.n., is morphologically distinct from typical S. tenuissima and represents a different species, which is assigned herein to the newly described S. monticola .

Figures

FIGURE 2. SEM micrographs of stem section and leaves of Selaginella tenuissima (Brade 9662, R). A. Upper surface of stem. B. Upper surface of median leaves. C. Close-up of apex of median leaf. D. Close-up of distal portion of inner half of median leaf lamina, upper surface. E. Lower surfaces of lateral leaf and of proximal portion of median leaf outer half; note marginal stomata (a). F. Close-up of proximal portion and margin of acroscopic half of lateral leaf lamina, lower surface.

FIGURE 3. Lectotype specimen of Selaginella tenuissima (Glaziou 4499, P [P00573924]). A. Close-up of characteristic label of Glaziou’s herbarium with his handwriting, including specific information on type locality as in protologue. Original image provided by and reproduced with permission of the vascular plants collections (P) of the Muséum national d’Histoire naturelle, Paris (France).

FIGURE 4. Image of specimen of Selaginella tenuissima (Glaziou 4499, P [P00044826]); note label lacking specific information included in protologue of the species. A. Close-up of upper surface of stem. B. Close-up of median leaves, upper surface. C. Close-up of strobilus; note white megaspore and sporophylls. Original images provided by and reproduced with permission of the vascular plants collections (P) of the Muséum national d’Histoire naturelle, Paris (France).

FIGURE 5.Image of specimen of Selaginella tenuissima (Glaziou 4499, C.[C10020554]); note labellacking specific information included in protologue of the species. Original image provided by and reproduced with permission of the Natural History Museum of Denmark.